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Probainognathus
Probainognathus (Greek: "progressive jaw") is a genus of meat-eating mammal-like reptile (Therapsids) that lived during the lower Upper Triassic of South America. This creature had an incipient squamosal-dentary jaw-cranium joint, which is a clearly mammalian anatomical feature. It was at the very least closely related to the family of Chiniquodontidae, in which some authors have included it. Some broadly similar teeth from Europe were described under the name of Lepagia. Known from about three dozen specimens, this creature was only about 10 cm long. Two skulls, including the holotype, were stolen from the University of La Rioja, Argentina, in February 1994. A pair of Probelesodon holotypes, (P. lewisi and P. minor), accompanied them. Etymology Probainognathus was first described and named by Harvard paleontologist, Alfred Sherwood Romer in 1970.1 Probainognathus jenseni is the focus of this description, and was named generically for its advanced jaw articulation, literally meaning “progressive jaw” in Greek, and specifically for James A. Jenson, a Harvard collector on the trips to recover the specimen. Description and paleobiology Probainognathus was a small cynodont, the apparent body size of which is inconsistent across sources, ranging from being about the size of a rat, to the size of a small dog. Teeth and diet The diet of Probainognathus is thought to have been carnivorous, which can be extrapolated by its teeth.1 The teeth include dully-tipped, partially developed canines, postcanines, four upper incisors that are positioned vertically, and three lower incisors that lean slightly forward.7 The cheek teeth are slender from the sides, but are lengthened anteroposteriorly, with several cusps along the row.1 This pattern is consistent with that exhibited by other meat-eating cynodonts, which indicates Probainognathus was a carnivore, as well. This tooth pattern is similar to that of chiniquodontids, and is part of the proposed connection between them and Probainognathus. Brain Based on an endocranial cast of Probainognathus, several details have been extrapolated about its brain. Probainognathus’ cerebral hemispheres were elongated, it had established olfactory bulbs, and a significant cerebellum and flooculi.9 These were among other, less pronounced, structures such as anterior colliculi.9 Because of these features, it has been suggested that Probainognathus’ brain had visual, auditory, motor, and body sensory systems all present. And, as a whole, the brain of Probainognathus indicates that it was in the process of developing endothermy. Skull The snout of Probainognathus extends beyond the premaxillary processes, and is very thin. The apparent fragility of this structure has led to it being broken off several specimens, and has led to an increased difficulty in studying it.1 There are several features that have been identified, however. The suborbital arch and the zygomatic arch are both shaped relatively sharply, and there is an extension of the squamosal posteriorly on the zygomatic arch. Probainognathus has a long secondary palate as it stretches all the way to the posterior end of the tooth row.1 In addition, the secondary palate is also formed largely by the palatine, and it is somewhat narrow. This narrowing of the palate allows for the conservation of space to be able to fit the back, lower molars. Moreover, the vomer is posterior to the secondary palate, and significant pterygoid flanges are present, as well.1 The skull of Probainognathus is, on average, around 7 cm in length, and is missing the parietal foramen.1 The skull is broad in proportion to its length, the face is short and narrow, but its zygomatic region is relatively large, seemingly for jaw muscle attachment.1 The skull also possesses a double occipital condyle, which is a typical mammalian feature.8 In comparison to close relatives, such as Probelesodon, the braincase of Probainognathus is more broad in the parietal region, and the oticoccipital region of the skull is deeper. Likewise, the parietals have more depth, the occipital crest is higher, and the orbits are proportionately larger relative to the rest of its head. The sagittal crest is also lengthened posteriorly, and splits into the occipital crests. This leads to a straighter posterior margin, and a more concave occiput in Probainognathus. Jaw The jaw of Probainognathus is of particular phylogenetic importance. Morphologically, the dentary makes up most the lower jaw, and it curves and extends down posteriorly to the area of the articular and jaw articulation.1 Correspondingly, in the upper jaw, the squamosal bone becomes situated next to the quadrate.4 The posterior end of this enlarged dentary fits into a small nook in the squamosal of the upper jaw, and displays the beginning of the evolution of the squamosal-dentary jaw joint.14 This squamosal-dentary jaw articulation is the same joint exhibited by mammals, and the inclusion of this joint in Probainognathus indicates that it is a precursor to mammals. Along with this osteological feature, the jaw of Probainognathus is suggested to have had mammal-like soft tissue advancements, as well. The jaw muscles in Probainognathus were thought to have been positioned farther forward, with the masseter splitting into two separate muscles: the superficial masseter and the deep masseter.1011 Despite this new development, the reptilian quadrate-articular jaw joint persists in Probainognathus, and a paired jaw joint is the result. Because of this, the jaw of Probainognathus remains distinct from that of mammals due mostly to the presence of the articular and the quadrate.4 Once the dentary-squamosal articulation becomes more established, the former bones involved in jaw articulation, the articular and quadrate, can become integrated into the inner ear as the malleus and incus, respectively.1 This has not yet happened in the case of Probainognathus, but the reduced size of the quadrate, as well as its loose association with the squamosal and proximity to the stapes indicates the quadrate to incus prosses is underway.8 This combination of evidence further solidifies Probainognathus’ phylogenic placement on the line to Mammalia, and provides a sound evolutionary connection between reptiles and mammals. Geological and paleoenvironmental information Category:Cynodonts Category:Triassic synapsids Category:Synapsid stubs Category:Prehistoric synapsids of South America